Lee & Thomas: Taxonomic Status of clavipes 145

CLARIFICATION OF THE TAXONOMIC STATUS OF CUCUJUS CLAVIPES WITH DESCRIPTIONS OF THE LARVAE OF C. C. CLAVIPES AND C. C. PUNICEUS (COLEOPTERA: )

JONGEUN LEE1 AND MICHAEL C. THOMAS2 1Department of Biological Science, College of Natural Sciences, Andong National University, Andong, 760749 E-mail: Korea [email protected]

2Florida State Collection of , Division of Plant Industry, Gainesville, Florida 326147100, USA E-mail: michael.thomas@freshfromflorida.com

ABSTRACT

The larvae of Cucujus c. clavipes Fabricius and C. c. puniceus Mannerheim are fully de- scribed and illustrated in detail for the first time. Based on larval and adult morphology the present recognition of two subspecies is maintained.

Key Words: , Cucujus, larva, North America

RESUMEN

Por primera vez se describen e ilustran las larvas de Cucujus c. clavipes Fabricius y C. c. pu- niceus Mannerheim. Basándose en la morfología larval, se acepta el reconocimiento de las dos subespecies.

Translation provided by the authors.

Cucujus clavipes Fabricius (1781) was de- (1931) described Cucujus clavipes subnitens as a scribed from “America boreali.” Cucujus pu- variety from Arizona and Utah. Thomas (1993) in a niceus Mannerheim (1843) was described from list of Nearctic Cucujidae treated C. puniceus as a “insula Sitkha”, now Baranof Island in south- subspecies of C. clavipes and Schaeffer’s taxon as a eastern Alaska and the site of the modern city variety as previously described. of Sitka. Both descriptions are of adults only, In an effort to resolve the status of Cucujus are based on the adult stage and are brief and clavipes we examined adults and larvae from both relatively uninformative. Of C. clavipes, Fabri- eastern and western North America. cius wrote: “ruber, thorace fuscato, femoribus clavatis rufis” (red, thorax dark, femora clavate, Larvae red); of C. puniceus, Mannerheim wrote: “elong- atus, depressus, laete sanguineus, antennis ni- Larvae of Japanese Cucujus coccinatus grofuscis, pectore abdomineque rufoferrugineis, Lewis were described and illustrated by Ha- thorax subrotundato, lateribus leviter denticu- yashi (1980, 1986) and the larva of C. mniszechi lato, supra obsolete bisulcato” (Elongate, de- Grouvelle was described by Lee and Sato pressed, rich red, antennae nigro-fuscus, abdo- (2007). men rufo-ferrugineous; thorax rounded, later- Larvae of C. clavipes Fabricius were briefly and ally weakly denticulate, above obsoletely bisul- partially illustrated (head and mandible) by Bøv- cate). ing and Craighead (1931) and Klausnitzer (2001). LeConte (1854, 1861, 1863) consistently treated Peterson (1951) provided extensive illustrations of C. puniceus as a valid species. Casey (1884) re- C. clavipes but provided only a brief description. In duced it to a variety of C. clavipes and said of it: neither case was the origin of the specimen illus- “The body is more elongated, and usually of a trated provided. Lawrence (1991) re-used Peter- brighter color. The first joint of the antennae is son’s illustrations and added scanning electron mi- usually of a dark testaceous, while in clavipes it is crographs of mouthparts of a specimen from Cali- black. The antennae are slightly longer, and the fornia. The larva of both North American subspe- neck slightly narrower in puniceus.” Leng (1920) cies of C. clavipes Fabricius are fully illustrated treated C. puniceus as either a variety or subspe- and described for the first time in the present pa- cies of C. clavipes [In the Leng Catalogue, a let- per. The larva of C. clavipes is similar to C. mnisze- tered taxon following a numbered species name chi (Lee and Sato 2007), but can be distinguished could be “. . . variety, subspecies, race, etc.” (Leng by absence of a distinct epicranial stem and pres- 1920: v)] and Hetschko (1930) followed Casey in ence of a sharp prostheca. In C. mniszechi the epi- treating it as a variety of C. clavipes. Schaeffer cranial stem is present and the prostheca is blunt.

146 Florida Entomologist 94(2) June 2011

Larvae of C. clavipes are reported to be preda- suture complete. Labrum (Fig. 1G) free, with 3 ceous (Smith and Sears 1982) or facultatively pre- pairs of setae and anterior border fimbriate. daceous (Lawrence 1991). Their extreme cold tol- Epipharynx glabrous medially, with 5 anterior se- erance, which increases with increasing latitude, tae on each side. Antennae 3segmented, ratio of has been extensively studied (Sformo et al. 2010, lengths of antennomeres 1, 2, and 3 about 1.0: 1.2: and references therein). 1.0. Mandibles (Fig. 1H) heavily sclerotized, sym- metrical, apices bidentate with a smaller subapi- MATERIALS AND METHODS cal tooth; with 2 dorsolateral mandibular setae; prostheca acuminate, spinelike, with a broad The larvae were preserved in 70% ethyl alco- base; mola with numerous setae medially and hol, cleared in 10% KOH solution for 1 hour, penicillus posteriorly (The scanning electron mi- rinsed in water, and dissected under a stereo- crographs in Lawrence (1991: 464, figs. 34.528, c- scopic microscope (Leica® MS5). Slide mounting f) show a conspicuous patch of microtrichia on procedures were carried out according to LeSage both the dorsal and ventral surfaces of the man- (1984), and the larval terminology follows dible near the base; these are virtually invisible Lawrence (1991). Specimens were measured with in liquid and are not illustrated here). Maxilla an ocular micrometer and the measurements (Fig. 1E) with cardo triangular, divided by an in- were transferred to graph paper. The illustrations ternal ridge, basal portion trapezoidal, 1 moder- were then sketched in pencil, the sketches inked, ately elongate seta near latero-basal margin; sti- and assembled into plates, which were optically pes elongate; mala falciform with 5 apical spines scanned and cleaned up in a graphics editor. Spec- and a medial brush composed of several thick se- imens examined are deposited in the Florida tae; maxillary palpus 3segmented, segment 1 ase- State Collection of Arthropods (FSCA) and the tose, segment 2 with 2 setae, segment 3 with 4 University of Alberta E. H. Strickland Entomo- minute apical setae. Labium (Fig. 1F) with con- logical Museum (UASM). spicuous mentum and prementum; mentum about as long as wide, with 2 pairs of setae and Descriptions prementum with 1 pair of setae and 1 pair of sen- silla; ligula rounded anteriorly, 1 pair of setae and microtrichia anteriorly; labial palpi 2-segmented Cucujus clavipes clavipes Fabricius, 1781 (Fig. 1 AJ) and widely separated at base. Thorax: Meso and metathorax tergites, and Diagnosis: See this section under C. c. puniceus. abdominal tergites and ventrites 18 each with 1 transverse ridge near anterior margin, ridge on Material examined: 37 total from: INDIANA: ventral surface of abdominal segment 1 lightly Morgan Co.: Martinsville (10); Tippecanoe Co. (1); sclerotized. Prothorax subquadrate, transverse, OHIO: Champaign Co. (1); Columbiana Co. (1); 0.5 times as long as wide, sides slightly curved, WISCONSIN: Calumet Co.: Forest Junction (1); dorsal surface smooth; prosternal surface smooth, Ingham Co.: Dansville State Game Area (1); Sha- 3 setae (1 elongate) at anterolateral angles and 2 wano Co.: Shawano (16); Shawnee Co.: Tilleda (6) short setae at posterolateral angles; prosternum (all deposited in the FSCA). trapezoidal, sides oblique, posterior margin Description: Late instar (Fig. 1A). Body 22.0 - straight, pair of medial setae present posterior to 26.0 mm long, elongate, subparallel, strongly dor- posterior margin of presternum. Meso- and met- soventrally flattened with strongly forked median athorax transverse, both 0.5 times as long as process at abdominal apex (Fig. 1A). Head and wide, sides curved, dorsal surface of both tergites abdominal segment 8 moderately sclerotized, yel- smooth with 3 short setae at anterolateral angles lowishbrown to brown, tergite of abdominal seg- and 2 short setae at posterolateral angles; both ment 9 strongly sclerotized and brown. sterna without well-defined subdivisions, each Head (Fig. 1B): prognathous, strongly trans- smooth with a pair of discal setae near anterior verse and dorsoventrally flattened. Lateral mar- margin; spiracular sclerite projecting strongly gin rounded. Median endocarina absent; epicra- from lateral margin, spiracles (Fig. 1C) annular nial stem present but very short; frontal sutures and angled posterolaterally. Legs (Fig. 1D) mod- lyriform, strongly curved; bases contiguous. St- erately long, 5segmented; claw falciform, large. emmata well-developed, 6 on each side of head Abdomen: Segments 17 transverse, tergite (Peterson (1951) reported 5 on each side; we count surface smooth with 2 setae anterior to spiracles 6 but 1 is small and difficult to see). Frontoclypeal and 2 setae posterior to spiracles; ventrite surface suture absent. Fronotoclypeal region with 3 long with 3 setae, 2 anteriorly and 1 posteriorly. Seg- setae anterior to angles of frontal arms, 1 pair an- ment 8 slightly enlarged, tergite (Fig. 1I) with a terior to the apex of the frontal arms on each side stout spicule at each posterolateral margin, pos- of the head, 1 pair medially between the frontal terolateral angles with 4 long and 4 short setae, 3 arms, and 1 pair at the apex of the frontoclypeal pairs of short setae anteromedially; sternite (Fig. region near the clypeolabral suture. Clypeolabral 1J) with 7 pairs of setae and with large stout pro-

Lee & Thomas: Taxonomic Status of Cucujus clavipes 147

Fig. 1. Larva of Cucujus c. clavipes. A, habitus, dorsal view; B, head, dorsal view; C, A7 spiracle, D, prothoracic leg; E, left maxilla, dorsal view; F, labium, ventral view; G, labrum, dorsal view; H, left mandible, dorsal view; I, ab- dominal segments 89, dorsal view; J, same, ventral view.

148 Florida Entomologist 94(2) June 2011 cess posteriorly with many minute setae apically. segment 1 asetose, segment 2 with 3 setae, seg- Tergum 9 with a basally forked process, directed ment 3 with 1 seta and 4 minute apical setae. La- dorsad; base of process with a pair of short, api- bium (Fig. 2F) with conspicuous mentum and pre- cally forked processes, 1 short seta at apex of mentum; mentum about as long as wide, with 3 forked process; anterior margin with laterally pairs of setae, prementum with 3 pairs of setae; curved processes projecting from tergum 8; ven- ligula transverse, with anterior microtrichia; la- trite 9 reduced and concealed from above. bial palpi 2 segmented. Thorax: Meso and metathorax tergites, and Cucujus clavipes puniceus Mannerheim abdominal tergites and ventrites 18 each with 1 (Fig. 2 AJ) transverse ridge near anterior margin, ridge on ventral surface of abdominal segment 1 smaller Diagnosis. The larva of this species is very sim- lightly sclerotized. Prothorax subquadrate, trans- ilar to that of Cucujus c. clavipes, but can be dis- verse, 0.5 times as long as wide, sides curved, dor- tinguished by the ratio of the 8th abdominal seg- sal surface smooth; prosternal surface smooth, 3 ment length vs length of the forked process (4:3 in setae (1 elongate) at anterolateral angles and 2 C. c. puniceus; 1:1 in C. c. Clavipes), and the ratio short setae at posterolateral angles; prosternum of the 8th abdominal segment width vs the width trapezoidal, sides oblique, posterior margin of forked process (measured at tips) (5:3 in C. c. straight, a pair of medial setae present posterior puniceus; 3:2 in C. c. clavipes). to posterior margin of presternum. Meso- and Material examined: 7 total, from: CANADA: metathorax transverse, both 0.5 times as long as ALBERTA: George Lake (2, UASM); USA: CALI- wide, sides curved, surface of both tergites FORNIA: El Dorado Co.: Blodgett Forest (1, smooth with 3 short seta at anterolateral angles FSCA); Tulare Co.: Sequoia National Park, and 2 short setae at posterolateral angles; both Stoney Cr. Picnic Area (2, FSCA);UTAH: Cache sterna without well- defined subdivisions, each Co.: Logan Valley (2, FSCA) smooth with a pair of discal setae near anterior Description: Late instar larva (Fig. 2A). Body margin; spiracular sclerite projecting strongly 21.0-24.0 mm long, elongate, subparallel, from lateral margin, spiracles (Fig. 2C) annular strongly dorsoventrally flattened with forked me- and angled posterolaterally. Legs (Fig. 2D) mod- dian process at abdominal apex (Fig. 2A). Head erately long, 5segmented; claw falciform, with 2 and abdominal segment 8 moderately sclerotized, setae. brown, tergum 9 strongly sclerotized and dark Abdomen: Segments 17 transverse, tergite brown. surface smooth with 2 setae anterior to spiracles Head (Fig. 2B): prognathous, strongly trans- and 2 setae posterior to spiracles; ventrite surface verse and dorsoventrally flattened. Lateral mar- with 3 setae, 2 anteriorly and 1 posteriorly. Seg- gin rounded. Hind corners of epicranium slightly ment 8 enlarged, tergite (Fig. 2I) with a stout sp- produced posteriorly. Median endocarina and epi- icule at each posterolateral margin, posterolat- cranial stem very short; frontal sutures lyriform, eral angles with 8 short setae, 3 pairs of short se- strongly curved; bases contiguous. Stemmata tae anteromedially, 2 pairs of short setae postero- well-developed, 6 present on each side of head. Fr- medially. Ventrite (Fig. 2J) with 9 pairs of setae ontoclypeal suture absent. Fronotoclypeal region and large stout process posteriorly with numer- with 3 long setae anterior to angles of frontal ous minute setae apically. Tergite 9 with a basally arms, 1 pair anterior to the apex of the frontal forked process, directed dorsad, as wide as long; arms on each side of the head, 1 pair medially be- base of process with a pair of short, apically tween the frontal arms, and 1 pair at the apex of forked processes, 1 short seta at apex of forked the frontoclypeal region near the clypeolabral su- process; anterior margin with lateral curved pro- ture. Clypeolabral suture complete. Labrum free cesses projecting from tergite 8; sternite 9 re- (Fig. 2G), with 5 pairs of setae. Epipharynx medi- duced and concealed from above. ally glabrous, 6 anterior setae on each side. An- tennae 3segmented, ratio of lengths of antenno- Adults meres 1, 2, and 3 about 1.0: 1.4: 1.0. Mandibles (Fig. 2H) heavily sclerotized, symmetrical, apices Given the differences discovered in the larvae bidentate with a smaller subapical tooth; with 2 of the 2 subspecies, we examined adults to deter- dorsolateral mandibular setae present; prostheca mine if there were corresponding adult differ- acuminate, spinelike, with a broad base; mola ences. We examined 120 adult specimens of C. c. with numerous setae medially and posteriorly. clavipes in the FSCA from the following states Maxilla (Fig. 2E) with cardo, divided by an inter- and provinces: CANADA: Ontario; USA: Colo- nal ridge, basal portion trapezoidal, with 1 mod- rado, Illinois, Indiana, Iowa, Kansas, Maine, erately elongate seta near basal margin; stipes Maryland, Massachusetts, Michigan, Mississippi, elongate; mala falciform, mala falciform with 5 Missouri, New York, New Jersey, North Carolina, apical spines and a medial brush composed of sev- Ohio, Pennsylvania, Virginia, Wisconsin. We ex- eral thick setae; maxillary palpus 3segmented, amined 46 adult specimens of C. c. puniceus in the

Lee & Thomas: Taxonomic Status of Cucujus clavipes 149

Fig. 2. Larva of Cucujus c. puniceus. A, habitus, dorsal view; B, head, dorsal view; C, A7 spiracle, D, prothoracic leg; E, left maxilla, dorsal view; F, labium, ventral view; G, labrum, dorsal view; H, left mandible, dorsal view; I, ab- dominal segments 89, dorsal view; J, same, ventral view.

150 Florida Entomologist 94(2) June 2011

FSCA from the following states and provinces: bus, descriptionibus. Tom. 1. Hamburgi et Kilonii, CANADA: Alberta, British Columbia; USA: impensis C. E. Bohnii, 566 p. Alaska, California, Idaho, Oregon. HAYASHI, N. 1980. Illustrations for identification of lar- As noted in previous literature, C. c. clavipes vae of (Coleoptera) found living in dead has a black scape, while C. c. puniceus has a red trees in Japan. Mem. Education Inst. Private Schools Japan. No. 72, 53 pp. scape. However, specimens of C. c. puniceus from HAYASHI, N. 1986. 113 plates of Coleopteran larvae, In Alaska have black scapes. We had formed the im- K. Morimoto and N. Hayashi [eds.], The Coleoptera pression that individuals from the western U.S. of Japan in Color, Vol. 1, Osaka: Hoikusha, vi+323 were on average more elongate than those from pp., 113 pls. the eastern part of the country. Measurements of HETSCHKO, A. 1930. Cucujidae. Coleopterorum Catalo- series from both populations revealed consider- gus 15(109): 193. able overlap in body proportions, with specimens KLAUSNITZER, B. 2001. Die Larven Der Kafer Mitteleu- of the C. c. puniceus slightly more elongate, rang- ropas (6. Band ). Spektrum Akad. Verlag ing in size from 12.5mm to 16.6mm, while speci- Heidelberg, Berlin, 1301. LAWRENCE, J. F. 1991. Cucujidae. pp. 463488 In F. W. mens of C. c. clavipes ranged in size from 9.5mm Stehr, [ed.], Immature . Vol. 2. Dubuque, IA: to 14.6mm. Kendall/Hunt. Lee and Sato (2007) found taxonomically use- LECONTE, J. L. 1854. Synopsis of the Cucujides of the ful genitalic differences among Asian species of United States. Proc. Acad. Nat. Sci. Philadelphia 7: Cucujus. Male genitalia from specimens of C. 7379. clavipes from all parts of its distribution were ex- LECONTE, J. L. 1861. Classification of the Coleoptera of amined and found to be indistinguishable. North America. Prepared for the Smithsonian Insti- tution. Part 1. Smithsonian Misc. Collection 136: 1208. CONCLUSIONS LECONTE, J. L. 1863. List of the Coleoptera of North America. Prepared for the Smithsonian Institution. Despite the larval differences, the lack of con- Part 1. Smithsonian Misc. Collection 140: 149. sistent and significant morphological differences LEE, C. F., AND SATO, M. 2007. A review of the in the adults suggests that at this point given the Cucujus Fabricius (Insecta: Cucujoidea: Cucujidae) state of our knowledge, the present treatment of from Taiwan, Japan, and China, with descriptions of these 2 populations as subspecies of the same spe- two new species and the larvae of Cucujus mniszechi cies is valid. Research into molecular differences Grouvelle. Zool. Studies 46: 311321. LENG, C. W. 1920. Catalogue of the Coleoptera of Amer- may prove useful in understanding the limits of ica, North of Mexico. Mount Vernon, NY. John D. both taxa. Sherman, Jr. x + 470 pp. LESAGE, L. 1984. Immature stages of Canadian Neochlamisus Karren (Col.: Chrysomelidae). Can. ACKNOWLEDGMENTS Entomol. 116: 383409. We thank Chi Feng Lee and John Marris and 2 anon- PETERSON, A. 1951. Larvae of Insects. An Introduction ymous reviewers for reviewing a previous draft of this to Nearctic Species. Part II. Coleoptera, Diptera, manuscript. George Ball generously lent larvae from Neuroptera, Siphonaptera, Mecoptera, Trichoptera. the University of Alberta collection. This is Entomology Columbus, OH: A. Peterson. v + 435 pp. Contribution No. 1180 of the Bureau of Entomology, SCHAEFFER, C. 1931. On a few new and known Co- Nematology, and Plant Pathology, Florida Department leoptera. Bull. Broooklyn Entomol. Soc. 26: 174176. of Agriculture and Consumer Services. This research SFORMO, T., WALTERS, K., JEANNET, K., WOWK, B., was supported by a grant to the senior author from the FAHY, G. M., BARNES, B. M., AND DUMAN, J. G. 2010. 2008 Academic Exchange Program of Andong National Deep supercooling, vitrification and limited survival University. to -100°C in the Alaskan Cucujus clavipes pu- niceus (Coleoptera: Cucujidae) larvae. J. Exper. Biol. 213: 502-509 REFERENCES CITED SMITH, D. B., AND SEARS, M. K. 1982. Mandibular struc- ture and feeding habits of three morphologically BØVING, A. G., AND CRAIGHEAD, F. C. 1931. An illustrat- similar coleopterous larvae Cucujus clavipes (Cu- ed synopsis of the principal larval forms of the order cujidae), Dendroides canadensis (Pyrochroidae), and Coleoptera. Entomol. Am. (New Series) 11: 1351. Pytho depressus (). Can. Entomol. 114: CASEY, T. L. 1884. Revision of the Cucujidae of America 173175. North of Mexico. Trans. Am. Entomol. Soc. 11:69112. THOMAS, M. C. 1993. The flat bark of Florida FABRICIUS, J. C. 1781. Species insectorum, exhibentes (, , ). Ar- eorum differentias specificas, synonyma auctorum, thropods of Florida and Neighboring Land Areas 15: loca natalia, metamorphosin, adjectis observationi- i-viii and 1-93.